Biodiversity in Planted Forests - Forestry Commission

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Andrew Peace. 1. 2. 3. 4 ..... Robin Sturdy, Phil Ratcliffe, Bill Mason, Graham. Pyatt, Simon Hodge ... Ferris-Kaan, R., Humphrey, J.W. and Peace, A.J.. (1998).
Biodiversity in Planted Forests by Jonathan Humphrey, Richard Ferris, Martin Jukes and Andrew Peace

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Overmature Sitka spruce: a natural reserve to encourage structural diversity and deadwood habitats

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Long-tailed tit nesting in gorse within a forest ride

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Chickweed wintergreen, a characteristic herb of northern forests

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Hoverflies feeding on leopard’s bane

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Introduction

structure, and the occurrence of particular tree

The majority of planted forests in Great Britain were

link between such indicators and wider biodiversity

established in the 20th century, usually on previously

has not been substantiated to the same degree in

open ground with no recent history of forest cover

British forests as in other countries.

(Hodge et al., 1998), but occasionally through

In this article we present results from a 5-year

conversion of ancient semi-natural woodland

programme of research aimed at:

(Spencer and Kirby, 1992). Introduced conifer species such as Sitka spruce (Picea sitchensis) and Corsican pine (Pinus contorta var. maritima) make up a large proportion (66%) of the planted area. Opinions differ as to the potential value of these ‘new forests’ for biodiversity. Attention has often been drawn to deleterious effects on the flora and fauna of the habitats which forestry replaces or modifies (e.g. Ratcliffe and Thompson, 1989), but there have also been a number of studies which

species such as birch (Betula spp.). However, the

1. Obtaining baseline information on the types/levels of biodiversity in planted forests. 2. Evaluating the contribution of planted forests to the conservation of native flora and fauna through comparisons with semi-natural woodlands. 3. Identifying potential biodiversity indicators by relating the diversity of range of measured taxa to soil, climate, vegetation and stand structure variables.

have highlighted the positive value of planted forests

Summary data are presented for all taxa analysed to

for wildlife (e.g. Petty et al., 1995). However, these

date. Initial results from this programme were

studies have been mostly site specific, and there

presented in Forest Research Annual Report

have been no comparative studies of plantations of

1997–1998 (Ferris-Kaan et al., 1998). Ground

different crop species in contrasting bioclimatic

vegetation, fungi, lichen, bryophyte and selected

zones, or on a range of varied site types. Similarly,

invertebrate datasets are considered in more detail,

there have been few attempts to compare the

and potential indicators of diversity within these

biodiversity of planted forests with that of native or

groups are discussed. A number of management

semi-natural woodlands. With continuing pressures

options for improving habitat quality are identified.

on forest managers to improve the biodiversity of

A full analysis of the datasets for these and other

planted forests, e.g. the Forest Certification process

species groups are presented in a forthcoming

(Anon., 2000), baseline information is urgently

Technical Paper based on the proceedings of a

needed to provide a quantitative framework for

symposium held at Harrogate in November 2000

understanding the levels and types of biodiversity

(Ferris and Humphrey, in press).

currently found in plantations, and to inform the development of biodiversity enhancement strategies.

Study sites

An additional problem is that the comprehensive

Assessment sites were selected from within the

assessment of biodiversity is an extremely difficult

‘lowland’, ‘foothill’ and ‘upland’ bioclimatic zones of

task. It is rarely cost-effective or practical to

the Ecological Site Classification system (ESC; Pyatt

conduct a complete census of all taxa within a forest

and Suárez, 1997). The zones are defined by annual

stand, let alone an entire catchment or forest

precipitation totals (lowlands < 800 mm; foothills

landscape. Therefore, the identity of biodiversity

800–1500 mm; uplands >1500 mm). Study sites

‘indicators’ or surrogate measures of biodiversity

were established in the main crop and semi-natural

has become a research priority in recent years

woodland types within each zone (Figure 1). At the

(Ferris and Humphrey, 1999). Examples of potential

majority of sites, a chronosequence of four 1 ha

indicators include: deadwood, vertical stand

(100 m x 100 m) permanent sample plots was

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established in forest stands encompassing different

Practicality and cost also influenced the selection of

growth stages based on the normal economic

species groups to assess. Invertebrates were

rotation (1: pre-thicket, 2: mid-rotation, 3: mature, 4:

sampled from deadwood and three separate vertical

overmature). Wherever possible, individual

strata (canopy, sub-canopy, ground). Canopy

chronosequences were established on sites with

dwelling species were sampled using a canopy

similar soils, climate and site history.

insecticide fogging technique (Jukes et al., in press), sub-canopy species by malaise traps (Humphrey et al., 1999), and ground species by pitfall traps (Jukes et al., 2001). The frequency and abundance of

Key

Glen Affric Queens Clunes Beasdale Knapdale Taynish Glentress Kielder

Sitka spruce - uplands Sitka spruce - foothills

vascular plants and fungi were sampled in eight sub-

Oak - uplands Oak - lowlands

plots within each 1 ha plot (Ferris and Humphrey, in

Scots pine - foothills Scots pine - lowlands

press). Species frequency and abundance estimates

Corsican pine - lowlands Norway spruce - lowlands

were made for lichens and bryophytes growing on individual pieces of deadwood (Humphrey et al., in press). Volume and decay status of fallen deadwood (logs), standing deadwood (snags) and

Sherwood Fineshade

stumps were recorded within each plot. Songbird

Thetford

monitoring was undertaken by point counts and

Dean

territory mapping (Ferris and Humphrey, in press).

Windsor

F I G U R E

Alice Holt

Data analysis methods for the various groups are

New Forest

described in full in the publications cited above.

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Location of biodiversity assessment plots; 52 plots were sampled in total over a 4-year period.

Overview of results Over 2000 species have been recorded to date (Table 1) with nearly half of these being invertebrates. A surprisingly high number of species were recorded in the planted stands, with lowland Scots pine and Norway spruce stands being the

Assessment methods

most species-rich of all the crop types. Stands in northern Britain (foothills and upland) had generally

Plots were selected to minimise heterogeneity in

less diverse invertebrate and songbird communities

stand structure, species composition, topography

than those in the south, but richer lichen and

and hydrology. A standardised system of

bryophyte communities. It had been expected that

assessment was established to maximise potential

the native woodland stands would be considerably

comparisons between measured attributes and to

more species-rich than the plantations, but this was

minimise disturbance during sampling. The plots

only the case for some groups such as vascular

were permanently marked. Assessments were

plants and lichens. These results are due in part to

carried out over a 3–4 year period and covered:

under-sampling of some groups in the native stands

structural aspects of biodiversity, such as vertical

(e.g. birds, Table 1), and to a lower number of stands

foliage cover and deadwood; taxa important in

surveyed (e.g. 4 upland oak stands compared to 8

ecosystem functioning (e.g. fungi); a range of

upland Sitka spruce stands). However, the positive

different groups making up the ‘compositional’

value of plantations for groups such as the beetles

aspect of biodiversity (e.g. invertebrates).

and hoverflies is substantiated by other studies in

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Table 1 Total number of invertebrate, fungi, lichen, bryophyte, vascular plant and songbird species recorded in each forest/climate zone type. Ninety-four coleoptera species (from a total of 474 species records excluding carabids) were recorded in more than one of the three vertical strata (canopy, sub-canopy, ground). Lowland Corsican pine

Scots Norway pine spruce

Foothill Oak

Scots Sitka pine spruce

Upland Sitka spruce

Oak

Total species

Red Data species

Canopy invertebrates Coleopteraa

71

81

86

66

53

66

47

31

225

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Sub-canopy invertebrates Cicadomorphab Syrphidsc Coleopterad

55 27 76

68 43 109

72 37 114

– 15 80

35 25 61

22 22 52

33 29 52

– 22 62

133 59 228

0 4 4

Ground invertebrates Coleoptera (excluding carabids)e Carabidsf

36 30

54 21

52 29

25 16

30 18

29 23

35 17

19 18

116 53

1 1

Deadwood invertebratesg

23

24

21

8

20

14

23

2

64

3

Fungi Lichens Bryophytes Vascular plants Songbirds

94 11 25 28 18

249 29 35 34 25

170 14 32 47 27

181 51 37 55 –

210 100 31 27 17

88 23 39 29 16

232 46 54 40 15

127 102 60 60 –

677 202 111 143 40

29 2 0 0 0

494

772

701

534

627

423

623

503

2051

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Totals

a Beetles – 37 families; b cicalids – 5 families; c hoverflies; d beetles – 30 families; e beetles – 26 families; f ground beetles. g The deadwood invertebrates category comprises data from a number of insect groups, many of which remain to be analysed. It is anticipated that more species will emerge from the deadwood collected from the oak stands. – no data available.

planted forests (Humphrey et al., 1998). A scattering of Red Data list species were recorded

Ground vegetation

across the majority of species groups, but the most

Although ground vegetation diversity was closely

striking result was for the fungi where 29 Red Data

related to soil nutrient levels there was also an effect

list species were recorded, suggesting that

of stand age on vegetation community composition,

plantations provide a particularly valuable habitat for

with the vegetation in most crop types becoming

rare fungi. Fungi, however, have been under-

progressively more wooded in character through the

recorded in the past compared to other groups,

stand cycle (increase in woodland similarity

particularly in plantations, so many of these species

coefficient – Table 2). This process is a function of

may be less rare than previously thought.

stand age (enough time has elapsed to allow slowcolonising woodland plants to become established in the stand), but may also be related to site history, i.e. whether the stand was established on a site previously occupied by ancient semi-natural woodland. Nevertheless, the plantation coefficients are still lower than those of the semi-natural woodland ‘control’ plots (oak and overmature foothills Scots pine), and only time will tell whether the wooded character of the vegetation of the overmature planted stands will continue to develop.

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Table 2 Changes in the woodland vegetation similarity coefficient in relation to stand stage and crop type. The woodland vegetation similarity coefficient gives a measure of how closely the sampled vegetation is matched to a National Vegetation Classification (NVC; Rodwell, 1991) semi-natural woodland community type relative to a nonwoodland vegetation community. Woodland vegetation similarity coefficient (%)

Pre-thicket

Mid-rotation

Mature

Overmature

Lowland Corsican pine

43.2

32.8

27.4

58.2

Lowland Scots pine

21.9

34.6

34.9

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Foothills Scots pine

41.3

58.2

69.1

70.8

Lowland Norway spruce

56.6

41.9

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Foothills Sitka spruce

33.3

17.5

25.5

36.8

Upland Sitka spruce

44.7

33.9

49.9

58.6

Lowland oak



88.9

60.7



Upland oak



75.4

79.2



– Stand stage not available.

Invertebrates

Canopy Coleoptera (beetles) were categorised into

Carabid (ground beetle) species-richness and

predators and others. The main influences on

community composition were strongly influenced by

community composition were tree species and

latitude and canopy structure, and to a lesser extent

latitude. Saproxylic species showed an increase in

by soil organic matter and vegetation diversity.

abundance with plantation age, but no relationship

Species diversity was greatest in the more open

with fallen deadwood volume was found, possibly

plots, either pre-thicket or overmature, declining

due to the narrow sampling window (only one

with canopy closure. There was a significant

sample per year for 2 years in each plot). Predators

increase in the proportion of forest specialist species

were the most abundant group in many plots,

as the stands matured, with the overmature stands

particularly spruce, where the predators associated

having the highest values (Figure 2).

with the green spruce aphid, Elatobium abietinum,

the functional groups saproxyles, herbivores,

were dominant. Higher structural diversity in Scots

% of total carabid species

pine stands resulted in a more diverse canopy 50

Coleoptera community, though this relationship was

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not found in other tree species. A positive

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relationship between leaf area index and herbivore

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species richness in pine suggests that as stands are

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thinned, the loss of canopy cover has a direct effect on the species-richness of canopy Coleoptera.

0 pre-thicket

mid-rotation

mature

overmature

Large numbers of invertebrates, mainly Diptera

Chronosequence specialists

F I G U R E

generalists

others

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The increasing proportion of carabid forest specialists with plantation age, together with a corresponding decline in non-forest species.

(flies) and parasitic Hymenoptera (wasps) were collected from the malaise traps. Hoverflies (Syrphidae), tree hoppers (Cicadomorpha) and beetles (Coleoptera) were identified to species level. Analysis of these data is still in progress, but the key points which have emerged are:

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• Hopper and hoverfly species-richness and diversity

counts relating to denser canopy cover. This finding contrasts with those of other studies which have

decreases as mean tree basal area increases.

suggested that the fungal flora of conifer stands is • Beetle diversity is not related to mean basal area. • Pre-thicket plots have the highest hopper and

often less diverse than that of broadleaved stands (Villeneuve et al., 1989). It is possible that the denser stand conditions associated with spruce are

hoverfly diversity.

conducive to the development of mycorrhizal • Sitka spruce canopies support the lowest

communities by affording freedom from competing

diversity for all three insect groups.

ground vegetation and providing a higher tree root density for mycorrhizal associations.

Fungi

In the lowland plots, mycorrhizal species-richness was positively correlated with the number of

The fungi were broken down into four functional

potential host tree species (P < 0.05). This confirms

groups: mycorrhizals, parasites, litter saprotrophs

recent analyses indicating that many tree species in

and wood saprotrophs. The results shown in Figure

Britain are associated with distinctive

3a are for mycorrhizals (223 species in total) and

ectomycorrhizal fungi which, in some cases, may be

wood saprotrophs (180 species). Upland Sitka

host-specific (Newton and Haigh, 1998). Wood

spruce plots had the highest mycorrhizal species

saprotroph richness was strongly correlated (P < 0.05) with increases in fallen deadwood (log)

(a)

volume (Figure 3b), with the lowland Scots pine and Wood saprotrophs

Mean species count (no. ha-1)

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oak plots having the highest species numbers and

Mycorrhizas

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log volumes.

25 20 15

Bryophytes and lichens

10 5 0

There was little distinction between the bryophyte d an

la

w

pl

U

Lo

la

K

K

P

O

C

SS

S

N

O

nd

nd

nd

SP

SP

SS

lls hi

la

ot

w

Lo

Fo

w

Lo

d an

nd

lls hi

la

ot

w

pl

U

Lo

Fo

flora of planted stands and semi-natural woodland

(b)

(Table 1). Bryophyte species-richness was more closely related to crop type than climate, with

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Species count (no.ha-1)

spruce stands being richer than pine stands, regardless of climate zone. This observation has not

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previously been recorded in Britain, but is consistent 40

with Fennoscandian literature, where spruce is generally considered a more favourable habitat for

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bryophytes than pine (Esseen et al., 1997). Bryophytes were less affected by shading, and most

0 0

10

20

30

Volume (m3 ha-1)

F I G U R E

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spruce stands had a reasonable complement of species in all stand stages. The best stands appear to be those with high values for upper canopy cover

(a) Mycorrhizal and wood saprotroph species-richness in relation to crop type. SP: Scots pine, SS: Sitka spruce, NS: Norway spruce, CP: Corsican pine, OK: oak.

(i.e. mature and overmature stand stages). It is

(b) Relationship between wood saprotroph species counts and volume of fallen deadwood (lowland sites only).

and constancy of micro-climate.

possible that these stands offer a better combination of high humidity, adequate light levels

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upland and foothills stands than in the lowland stands (Table 1), substantiating existing views that oceanic conditions (and low pollution levels) in the north and west of Britain provide much better conditions for lichen growth (Rose, 1993). The seminatural pinewood and oak stands had richer lichen communities than the planted stands, relating to the

Mean species count (no. ha-1)

Lichen species-richness was much higher in the 60 50 40 30 20 10 0 pre-thicket

mid-rotation

occurrence of habitat features such as old trees,

mature

overmature

oak

Stand stage / crop type

shaded rocks and cliffs, and a long-continuity of

Uplands / foothills

Lowlands

woodland conditions (Plate 1). F I G U R E

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Lichen species-richness in relation to stand stage; oak is included for comparative purposes.

B

P L A T E

1

Ancient semi-natural Scots pine wood showing old trees and standing deadwood. (Simon Davey)

P L A T E

2

Typical group of pinewood lichens including Cladonia botrytes (B) on deadwood; Vaccinium berry added for scale. (Simon Davey)

Low light levels are considered to be highly detrimental to lichen growth (Rose, 1993). This

(Kruys et al., 1999) tend to support these findings.

explains why stand structure had such a significant

Larger and more highly decayed material appears to

effect on lichen species-richness, mid-rotation and

provide a better substrate for lower plant

mature stages having lower species counts than the

development (Ferris and Humphrey, in press).

pre-thicket and overmature stands (Figure 4). The

Snags in native pinewoods provide a key habitat for

pre-thicket stands are particularly important in the

the rare ‘pin-head’ lichen group (species of the

foothills Scots pine plots where stumps support

genera Calicales and Chaenotheca).

Cladonia species. A scarce and important species within the group is C. botrytes shown in Plate 2. Bryophyte species-richness was higher on logs and stumps, while snags were more important for lichens (Figure 5). Observations from overseas

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Mean number of species

3 2.5 2 1.5 1 0.5 0 logs

snags

stumps

Deadwood type

Bryophytes

Lichens

P L A T E F I G U R E

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Lichen and bryophyte species-richness on different types of deadwood.

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Deadwood is an important substrate for lower plants and fungi especially within natural reserves. (Simon Davey)

1. Maintenance of the patch-clearfelling system to

Conclusions and implications for management

ensure provision of early-successional habitat. 2. Extending rotations beyond normal economic felling age and establishing ‘natural’ (non-

The project has been successful in meeting its main objective of establishing a baseline dataset of the biodiversity of plantations across Britain. The most significant finding was that stands of non-native conifer species appear to provide suitable habitat for a wide range of native flora and fauna and should be viewed as making a positive contribution to

intervention) reserves. 3. Managing some stands under continuous cover regimes (sensu Mason et al., 1999). The latter two strategies would help to maintain and develop mature woodland conditions, including production of old trees, accumulation of deadwood (Plate 3) and diversification of the tree flora.

biodiversity conservation in the UK. There were considerable differences in value of different stand

While wind risk is a significant factor to consider

ages and crop types for different species groups

when deciding where to locate continuous cover

with no single type providing ‘optimal’ conditions for

stands (Mason et al., 1999), site history is also of

biodiversity. Pre-thicket plots and restocks provide

key importance, as there is a suggestion from the

habitat for a variety of species-groups common to

data that plantations established on ancient semi-

non-wooded habitats (e.g. heathland). However, the

natural woodland sites (PAWS, e.g. some lowland

overmature stands, both in native and planted

Norway spruce and upland Sitka spruce stands) have

forests, are important for species requiring

a much higher diversity of woodland fungi,

continuity of wooded conditions and deadwood.

bryophytes and vascular plants than their counter-

Therefore, three contrasting, but not mutually

parts established on previously open ground.

exclusive, management strategies could be

Restoring PAWS back to native woodland may not

considered for plantations:

always be of benefit to biodiversity in the short to medium term, particularly if the restoration is done

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very rapidly. This is especially the case for

mycorrhizal diversity. The positive correlations

mycorrhizals where extensive felling of host trees

recorded between deadwood volumes, and species-

can disrupt mycelial connectivity and reduce the

richness of wood saprotroph fungi, lichens and

extent of recolonisation after restocking or natural

bryophytes is also notable. A mix of fresh and well-

regeneration (Flynn et al., 1998). Even smaller-scale

decayed large diameter logs and stumps should

felling can be disruptive (Flynn et al., 1998), and it

provide a range of habitat types (Plate 4) with

may be more desirable therefore to identify some

volumes in the range of about 40 m3 ha-1 providing

PAWS known to have a high number of

for the greatest lower plant and fungal diversity

characteristic species as potential ‘natural reserves’.

(Ferris and Humphrey, in press). The biodiversity indicators and management strategies proposed in this article are still tentative and further work is planned in the near future to validate these proposals and analyse additional datasets, looking particularly at the issue of site history.

Acknowledgements

The following individuals and organisations made a significant contribution to this research: Andrew Brunt, Ellie Dickson, Becky Lander, Su Meekins, P L A T E

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Old beech stump. Fresh and well-decayed large logs and stumps provide a range of habitat types. (Entomology Branch, Forest Research)

Antonia Nichol, Martin Schofield, Lorna Parker, Robin Sturdy, Phil Ratcliffe, Bill Mason, Graham Pyatt, Simon Hodge, Clive Carter, Derrick Hiscocks, Simon Davey, the Technical Support Unit of Forest Research, the British Mycological Society, the British

Based on the results presented above it is clear that measures of stand structure and deadwood have considerable potential as biodiversity indicators in planted forests. For example, the number of host tree species could be used as an indicator of

Trust for Ornithology, Forest Enterprise, Crown Estates, Highlands and Islands Enterprise, Alvie Estates. The research was funded by the Forestry Commission.

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